neoichnology of the seaward side of peixe lagoon in mostardas

Rev. bras. paleontol. 12(3):211-224, Setembro/Dezembro 2009
© 2009 by the Sociedade Brasileira de Paleontologia
doi:10.4072/rbp.2009.3.04
NEOICHNOLOGY OF THE SEAWARD SIDE OF PEIXE LAGOON IN
MOSTARDAS, SOUTHERNMOST BRAZIL: THE PSILONICHNUS
ICHNOCOENOSIS REVISITED
RENATA GUIMARÃES NETTO
PPGeo UNISINOS, Av. Unisinos, 950, 93022-000, São Leopoldo, RS. nettorg@unisinos.br
MARCELO ENGELKE GRANGEIRO
Deceased in June, 2005
ABSTRACT – This paper characterizes the modern Psilonichnus ichnocoenosis present in the seaward side of Peixe
Lagoon (southernmost Brazil) and discusses its preservation potential in the fossil record and its (paleo)environmental and
stratigraphic significances. This ichnofauna occurs in exposed softgrounds and stiffgrounds of the backshore and backbarrier
environments associated with the channel mouth area of the lagoon. The burrow assemblage is composed of Y- and J-shaped
dwelling burrows and trackways of grapsid crabs (Chasmagnatus granulata), fiddler crabs (Uca uruguayensis), and
ocypodid crabs (Ocypode quadrata), tracks and trackways of scarab beetles, birds, and small lizards, horizontal, randomly
branched, shallow feeding burrows of mole crickets, dwelling burrows of small rodents and ant nests. As a natural park,
most of the natural environmental conditions that support the establishment of such an ichnocoenosis are preserved in the
study area, favoring its use as a modern analogue to the ancient Psilonichnus Ichnofacies. Thus, original concepts developed
after the first record of the Psilonichnus ichnocoenosis in modern environments, which supported the construction of the
Psilonichnus Ichnofacies paradigm, are revisited herein and some premises are reinforced. Possible stratigraphic scenarios
are also estimated, allowing the inference that the best preservation of the Psilonichnus Ichnofacies occurs during the
establishment of transgressive system tracts.
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Key words: Psilonichnus ichnocoenosis, Psilonichnus Ichnofacies, crab burrows, marginal marine settings, Peixe Lagoon.
RESUMO – Este trabalho caracteriza a icnonocenose moderna de Psilonichnus presente na porção marinha da lagoa do
Peixe (extremo sul do Brasil) e discute seu potencial de preservação no registro fóssil e seu significado (paleo)ambiental e
estratigráfico. A icnofauna analisada ocorre em substratos moles e em substratos compactados plásticos de ambientes de
antepraia e atrás da barreira eólica associados à área de desembocadura do canal principal que alimenta a lagoa. A assembléia
é composta por trilhas e escavações de moradia em forma de Y e J de caranguejos (Chasmagnatus granulata, Uca uruguayensis
e Ocypode quadrata), pegadas e trilhas de aves, pequenos lagartos, besouros escarabeídeos, escavações horizontais, irregularmente ramificadas, rasas, feitas por grilotalpídeos em busca de alimento, galerias de pequenos roedores e ninhos de
formigas. Por ser um parque natural, as condições ambientais vigentes na lagoa do Peixe são muito próximas das de
ambientes intocados, o que favorece ao estudo da icnocenose de Psilonichnus ali presente como análogo moderno da
Icnofácies Psilonichnus. Assim, os conceitos originais, formulados a partir do primeiro registro da icnocenose de Psilonichnus
em ambientes modernos – e que serviram de base para a construção do paradigma que rege a Icnofácies Psilonichnus –, são
aqui revisitados e algumas premissas, reforçadas. A análise de possíveis cenários estratigráficos permite inferir que a melhor
preservação da Icnofácies Psilonichnus se dê durante o estabelecimento de tratos de sistemas transgressivos.
Palavras-chave: icnocenose de Psilonichnus, Icnofácies Psilonichnus, escavações de caranguejos, ambientes marginais
marinhos, lagoa do Peixe.
INTRODUCTION
coastal environments, such as those conducted by Hertwek
(1972), Howard & Dorjes (1972), Basan & Frey (1977), and
Frey & Pemberton (1987) on the Georgia coast (USA), Frey et
al. (1987) in the Yellow Sea tidal flats, Curran & White (1991),
White & Curran (1993), and Curran (1994) on San Salvador
Island (Bahamas), Gingras et al. (1999, 2000, 2002, 2004) in
Although being encouraged during the 1970s and 1980s,
neoichnological studies are still less extensive than those
focused on paleoichnological record. Also, most of the studies
on modern biogenic sedimentary structures are related to
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Willapa Bay (USA), De (2000) in the coastal Ganges Delta
Complex (India), Grangeiro & Netto (2003) in Peixe Lagoon
(southernmost Brazil), and Dashtgard & Gingras (2005) and
Hauck et al. (2009) on New Brunswick coast (Canada), among
others.
The Psilonichnus Ichnocoenosis, as defined by Frey &
Pemberton (1987), occupies supralittoral to upper littoral
substrates, deposited under moderate to low-energy marine
and (or) aeolian conditions, and subject to modification by
torrential rains or storm surges. Substrates are often
composed of well-sorted, variably laminated to crossstratified sand, to root- and burrow-mottled, poorly sorted
sand or muddy sand. They characterize coastal settings,
typically represented by beach backshore and dunes, but
also by washover fans, supratidal flats, and tidal inlets,
intergradational with the maritime terrestrial zone
(MacEachern, 2001). Vertical, irregularly J-, Y-, or U-shaped
shafts, some of them with bulbous basal cells, are the
predominant biogenic structures, representing ocypodid crab
or thalassinoidean shrimp-dwelling burrows, most of them
assigned to the Psilonichnus ichnogenus (Fürsich, 1981; Frey
et al., 1984; Nesbitt & Campbell, 2006). Invertebrate and
vertebrate crawling and foraging traces or superficial tunnels,
as well as microbial mats, vertebrate tracks and coprolites
can be locally present. The diversity of invertebrates is low,
being mostly predators or scavengers, while vertebrates could
be locally more diverse, being predominantly predators or
herbivores. In pre-Cretaceous units, crab-like dwelling
burrows may be absent (MacEachern, 2001).
Pemberton & Frey (1987) interpreted this ichnocoenosis
as representative of the Psilonichnus Ichnofacies, which
represents a mixture of marine, quasimarine and nonmarine
conditions. However, the common occurrence of Psilonichnus
upsilon in Pleistocene Bahamian backshore deposits
popularized the Psilonichnus Ichnofacies as a backshore
indicator (e.g. Curran & White, 1991; Gibert & Martinell, 1998;
MacEachern, 2001; Nesbitt & Campbell, 2006), although it
has a broader facies distribution in modern coastal settings.
Although Frey et al. (1990) had recognized the Psilonichnus
Ichnocoenosis as an archetypical ichnofacies, the non
observance of a long temporal distribution in geological
record supports the concern about its validity (Bromley &
Asgaard, 1991; Gibert & Martinell, 1998) and its spatial
distribution. Therefore, the aim of this paper is twofold: to
revisit the Psilonichnus Ichnocoenosis, based on its
occurrence in the seaward side of Peixe Lagoon, and to review
its distribution in the marginal marine settings, in light of the
predominant ecologic and sedimentologic aspects of the
study area.
producers, and understanding their behavior when interacting
with the substrate. As the study area is part of a national
park, a research license was required. The periodic visits
occurred every two months and lasted three days each time,
and the same study spots were systematically visited. The
biogenic sedimentary structures were observed, their external
morphology analyzed. The Y-shaped burrows made by
Chasmagnathus granulata were usually counted, and
differences in sizes and morphology of burrow entrances,
the number of individuals per burrow, sex and behavior were
considered, but no systematic statistics calculated. Data from
the lagoon benthic fauna and terrestrial biota living in or
around the lagoon settings, the chemical-physical aspects
of the lagoon and the surrounding substrates,
geomorphological and geographical characteristics,
sediments and sedimentary facies were also obtained and
analyzed.
Burrow architectures were accessed by casting, using
the uncolored acrylic resin RPI AROTEC. Water salinity was
systematically measured, from the lagoon and from the bottom
of the crab burrows, to access the salinity gradient and its
fluctuations in the study area. Biota were not collected, but
identified.
Sedimentological data (granulometry, presence/absence
of matrix, grain selection and mineral composition, physical
and biogenic sedimentary structures) acquisition consisted
of four SE/NW oriented transects which crosscut four NE/
SW oriented transects in the study area.
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MATERIAL AND METHODS
This study was based on periodic visits to the channel
mouth area of Peixe Lagoon between 1999 and 2002, and
sporadic visits from 2003 to 2008, with the objective of
analyzing the morphological and environmental aspects of
the present biogenic sedimentary structures, identifying their
GEOMORPHOLOGICAL AND SEDIMENTARY
SETTING
Geomorphological and sedimentary setting
Peixe Lagoon is a very shallow, 45-km-long water body
oriented NE-SW parallel to the coastline, between Mostardas and Tavares counties, in Rio Grande do Sul State,
southernmost Brazil (UTM N 6547000 and 6525000, and UTM
E 507000 and 485000, Central Meridian 51W; Figure 1). The
lagoon is formed by the waters that flow away from the two
main fluvial courses when they get to the depressed area, or
by marine waters, during storm surges. Its depth average is
0.5 m, except in the channels, where it can be as much as 4 m.
The barrier spit forming its seaward limit is approximately 100
m wide. Pleistocene sandstones and muddy sandstones of
the Lagoon-Barrier System III of Rio Grande do Sul Coastal
Plain (PCRS, Villwock et al., 1986) form the lagoon floor.
In the Peixe Lagoon area, PCRS overlies the Pre-Cambrian
granitic basement and the Tertiary deposits of Pelotas Basin.
To the east, it is bound by the modern aeolian sand dunes
and the sea (Figure 2). The west border is a small cliff (~ 5 m
high) formed by orange-yellowish quartz sandstones of
aeolian origin from the Barrier System III (Villwock, 1984).
The lagoon was formed after the last regressive event
that took place in PCRS, when the stream channels were
confined by a sand barrier, flooding the adjacent plains and
the surrounding shallow depressions (Tomazelli & Villwock,
2000, Figure 2). A constant supply of aeolian sediment source
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
Figure 1. Location map of the study area.
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Figure 2. Geological map of the Rio Grande do Sul Coastal Plain (PCRS), showing Peixe Lagoon confined between ancient deposits of
Barrier III and modern aeolian deposits of Barrier IV (modified from Tomazelli & Villwock, 2000).
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maintains the shallow depth of the lagoon. Sporadic storm
surges overwash the exposed sand barrier, introducing marine
waters into the lagoon and bringing marine organisms with
them. Most of these organisms return to the sea when the
water level falls, but some stay, forming orphan colonies.
During fairweather periods, the barrier is reconstructed and
the lagoon becomes confined again. The salinity, in general,
diminishes and only organisms tolerant to brackish-water
survive. Most of them are burrowers that rework the lagoon
floor and the adjacent marginal area. The thickness of water
laminae is reduced and a large amount of the lagoon floor is
exposed. The salinity gradient of the lagoon seaward-side
waters is around 5 to 15‰ (mesohaline), being always higher
than the inner zones, even when the barrier is up. The main
sedimentary deposits observed in the studied area are sandy
beaches, aeolian foredune ridge, discrete washover fans on
the backshore zone, washover channels, the lagoon floor, as
well as vegetated margins (Figure 3).
Beach deposits
Beach deposits are characterized by fine quartz sands
forming trough cross-stratification, accumulated as lenticular
bodies between the upper shoreface and the foreshore, up to
the seaward side of the backshore. Most of the foreshore
deposits are frequently exposed, as the tidal gradient is very
small (~ 0.5 m), being reworked by the wind, which forms
small eolic ripples, preserved chiefly in the transition zone to
the backshore. Longshore currents resulting from the
predominant NE-SW wind direction at Rio Grande do Sul
coast commonly rework the sands (Figure 3). Aeolian processes are dominant in subaerial conditions and construct a
sand bar in the channel discharge zone, isolating the lagoon
from the sea in fairweather periods. Storm surges usually
split the bar, generating discrete washover fans and
increasing the saline water input into the lagoon. Vertical
burrows of pelecypod Donax donax, which characterizes the
Skolithos ichnocoenosis in the study area, death marks of
jellyfish and trackways of isopod crustaceans are the most
frequent biogenic sedimentary structures in the swash line.
Ghost crab galleries (Ocypode quadrata) are the dominant
burrows in the transition zone to the backshore.
Aeolian foredune deposits
Yellow, well-sorted, bimodal fine sands forming the actual coastal barchanoid dune ridge and the associated interdune
zone, located in the backshore zone, parallel to the shoreline
and reaching 5 m in height (Figures 4A-C). Sand deposition
by the wind generates high-angle, tabular cross stratification
and sinuous or straight crest asymmetric ripples. The irregular evolution of the dunes produce avalanche sand tongues,
and adhesion ripples and tool marks are frequently observed.
Channels and ephemeral lakes are usually formed during the
storms, mobilizing the sands and creating small fans on the
seaward side of the lagoon, which develop into terraces.
Tracks and trackways of birds and lizards, horizontal,
branched, backfilled burrows of mole crickets (Orthoptera,
Grilotalpidae) and beetles (Coleoptera, Scarabeidae), and small
horizontal galleries of rodent mammals (Ctenomys ctenomys)
are the main biogenic sedimentary structures commonly
observed in dune and wet interdune deposits.
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Channel deposits
A main channel oriented NW-SE feeds the lagoon,
draining into the sea when the bar is open, being up to 4 m
deep and 200 m wide (Figures 4D-E). Sand is thrown out into
the channels by aeolian aspersion when the depth is less,
Figure 3. Satellite image from the Peixe Lagoon channel mouth area, showing the main settings in the studied area (image from Google
Earth, 2009).
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
building barrier islands. Neap tidal currents form straight crest
ripples and bidirectional cross stratification when the bar is
open. Biogenic sedimentary structures are observed in the
exposed channel margin deposits, which bear vertical Y-, Jand U-shaped burrows made by the amphibious crab
Chasmagnatus granulata (Y and U) and Uca uruguayensis
(J), as well as bird tracks and pelecypod vertical burrows
(Tagelus plebeus). Stable channel margins are vegetated by
grasses and mangroves.
Lagoon deposits
The flooded area adjacent to the channels characterizes
the lagoon deposits and is composed of organic rich muddy
sands (Figure 4F). The lagoon margin is periodically
expanded, exposing part of its floor, either due to the bar
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overflow due to storm surges, or due to low rainfall, in spring
and summer times. When exposed, muddy sand softgrounds
lose water and give rise to stiffgrounds or, seldom, firmgrounds
(Gingras et al., 2000, 2001). Biogenic sedimentary structures
inside the lagoon are produced by a low-diversity benthic
fauna, composed of mobile and stationary polychaetes, crabs
(Callinectes sapidus), pelecypods (T. plebeus), and
thalassinidean shrimp, which supply food to migratory birds.
Outside the lagoon, the edges are densely reworked by vertical, short Y- and J-burrows of respectively C. granulata
and U. uruguayensis. Tracks and trackways of crabs and
birds are also common and old openings of abandoned vertical shafts of polychaetes and T. plebeus can be seen when
the water laminae is very shallow, almost exposing the lagoon
floor.
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Figure 4. Main sedimentary deposits of the seaward side of Peixe Lagoon. A-C, Aeolian deposits, showing the barchanoid dune ridge
from Barrier IV (A), avalanche tongues and beetle burrows in the aeolian dunes (B), and the wet interdune deposits (C); D-E, channel
deposits, clearly delimited (D) or completely recovered during flooding periods (E); F, lagoon deposits, showing their shallowness and their
large and frequently exposed margins.
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THE PEIXE LAGOON PSILONICHNUS
ICHNOCOENOSIS
Two ichnocoenoses can be recognized in the seaward
side of Peixe Lagoon, one subaquatic and another subaerial
(Grangeiro & Netto, 2003). The subaquatic ichnocoenosis is
chiefly composed of vertical shafts and horizontal trails of
polychaetes, by shallow elliptical depressions made by crabs
and vertical V-shaped shafts of pelecypod siphons, when
the barrier is up and the lagoon is confined. The diversity of
subaquatic ichnocoenosis rises after storm surges, and stellar
grazing trails of polychaetes (?Nereis) and abundant
chimneys of thalassinid crustacean burrow systems can be
observed. This ichnocoenosis is representative of an
impoverished Cruziana Ichnofacies (sensu Pemberton &
Wightman, 1992).
The subaerial ichnocoenosis is dominantly composed of
modern Psilonichnus ichnogenus produced by amphibious
crabs. Invertebrate trackways and burrows, and vertebrate
tracks, trackways, and burrow systems also occur. The nature
of the main burrowers, the low ichnodiversity, and the
presence of biogenic sedimentary structures made by
scavengers (invertebrates), predators (invertebrates and
vertebrates) and herbivores (vertebrates) allow its
identification as a typical Psilonichnus Ichnocoenosis (Frey
& Pemberton, 1987; Pemberton et al., 1992a; Bromley, 1996;
MacEachern, 2001).
Five suites can be recognized in the Psilonichnus
Ichnocoenosis at Peixe Lagoon (Figure 5): (i) a lagoon edge
suite, which occurs in the backbarrier zone and is dominated
by short Y- and J-shaped burrows of C. granulata and U.
uruguayensis; (ii) a backshore suite, which occurs in the
beach side of the dune ridge and is dominated by large Jshaped burrows of O. quadrata; (iii) an aeolian sand dunes
suite, dominated by insect and vertebrate trackways; (iv) a
wet interdune suite, dominated by insect horizontal burrows;
and (v) a vegetated margin suite, dominated by multiple Ushaped burrows of C. granulata (Grangeiro & Netto, 2003).
The lagoon edge suite
This suite is composed essentially of burrows excavated
by mature adult crabs and shows short Y- and J-shaped
dwelling burrows made respectively by C. granulata and U.
uruguayensis. The J-shaped burrows of U. uruguayensis
(Figures 6A-B) are always excavated in the limiting zone of
the lagoon margins and the outer vegetated area, dominated
by grass, while the Y-shaped burrows of C. granulata (Figures 6C-E) are made in the bare stiffgrounds exposed between
the vegetated area and the water body. The Y-shaped burrows
often occur associated with thin vertical polychaete dwelling
burrows and, eventually, with discrete V-shape burrows of T.
plebeus, both belonging to the subaquatic ichnocoenosis.
Such association is, in fact, a palimpsest preservation that
records the seasonal oscillation of the lagoon water table
level. The crab burrows usually crosscut the polychaete and
pelecypod burrows, reflecting the dominance of periods of
low water table, when the lagoon edges expand. The J-shaped
burrows were occupied by the crab Metasesarma rubipres, a
clear evidence of opportunistic behavior.
The morphological features of the crab burrows observed
in the Peixe Lagoon margins allow to assignment as modern
Psilonichnus and to compare the Y-shaped burrows to
Psilonichnus upsilon. The depth of the main burrow observed
in the C. granulata galleries (33 cm long on average, Figure
6D) seems to be a direct response to the dominant high water
table level in the lagoon edge, which inundates the burrow’s
bottoms at a depth of 30 cm (see Figure 10), favoring branchial
respiration.
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Figure 5. Distribution of the Psilonichnus ichnocoenosis suites in the study area: 1, the lagoon edge suite; 2, the backshore suite; 3, the
aeolian dunes suite; 4, the wet interdune suite; 5, the vegetated margin suite.
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
Populations of C. granulata are more abundant than U.
uruguayensis, resulting in a 4:1 predominance of the Yshaped burrows. The crab populations follow the lagoon base
level fluctuations, colonizing the available moist substrate of
the seaward side during the whole year, but hiding
themselves in burrows during the winter. The Y-shaped
burrows are generally occupied by a couple and
extraordinarily by more than two crabs.
Avian tracks and trackways are other common biogenic
structures observed in the lagoon edge suite (Figure 6F).
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Most of the avifauna of Peixe Lagoon is migratory, which
stop over there to feed, during the whole year, or to breed,
chiefly in the southern hemisphere’s springtime. Trackways
are long and irregular, reflecting the animal’s movements, and
frequently accompanied by beak marks and open or broken
shells of T. plebeus, reflecting predation of these infaunal
bivalves.
The backshore suite
This suite is dominated by large inclined, J-shaped
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Figure 6. Biogenic sedimentary structures of the seaward side of Peixe Lagoon. A, J-shaped burrow of Uca uruguayensis with the
characteristic arrangement of pellets around the burrow entrance; B, cast of U. uruguayensis burrow; C, E, Y-shaped burrow of
Chasmagnathus granulata showing the scratch marks produced by the walking legs into and around the burrow entrance (C); D, cast of
C. granulata burrow; F, bird tracks and discrete bird trackways together with beak marks. Scale bars: 10 cm, with exception of B (10 mm).
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burrows of Ocypode quadrata (64 cm-long, on average, Figures 7A-C), which can also be assigned to the ichnogenus
Psilonichnus. O. quadrata burrows are longer than those
made by amphibious crabs in the lagoon edges and do not
show the Y-shaped openings. However, the main burrow
morphology is equivalent to the main burrow of the P. upsilon
ichnospecies. Burrows are sparsely distributed, occurring
preferentially close to the seaward side of the sand dunes.
Some few and discrete arthropod (insect and decapods) and
mammal trackways (rodents) are also observed in this suite
(Figures 7D, F).
The ichnogenus Psilonichnus has been attributed to crab
activity since the study of its significance carried out by
Frey et al. (1984). In spite of the reluctance of Fürsich (1981)
to attribute the type-species P. tubiformis to crab activity,
due to the incipient fossil record of these organisms in preCretaceous rocks, Frey et al. (1984) had no doubts to compare the Pleistocene Bahamian Psilonichnus upsilon with O.
quadrata burrows. The Bahamian P. upsilon occurs in beach
deposits, along the supratidal zone up to the dune barrier,
being attributed to the activity of adults of O. quadrata (Frey
et al., 1984; Curran & White, 1991). O. quadrata is broadly
distributed along the Atlantic coast from north to south
Hemisphere and Gulf of Mexico as well, reaching the Rio
Grande do Sul littoral (Buckup & Bond-Buckup, 1999). In
spite of Ocypode being the most “terrestrial” ocypodid
decapod, their burrows are confined to the zone between the
high-tide and the seaward side of the dune barrier. This fact
explains the occurrence of O. quadrata burrows only in the
backshore deposits in the study area, whilst the backbarrier
Psilonichnus were produced by grapsid and fiddler crabs.
collapsed in stiffgrounds, giving them a false bilobed aspect.
In softgrounds, the mole cricket burrows show a uniform
round and plain roof, with discrete external ornamentation,
such as small knobs. The burrows show meniscate infill,
resulting from the burrowing leg’s movement (Chamberlain,
1975), and always occur in wet and moist substrates.
Irregular trackways of dune scarab beetles and other
beetles are also common, composed of a continuous central
shallow depression with lateral ridges, formed by abdomen
dragging, and two rows of podia imprints, each one arranged
parallel to the ridges, in the outer part of the trackway.
Tetradactile bird tracks of distinct sizes forming straight
to curve trackways are the main vertebrate biogenic structures
in the wet interdune suite.
The vegetated margin suite
This suite is dominated by multiple U-shaped burrows of
C. granulata and by ant nests. The U-shaped burrows are
generally found in the lagoon margins stabilized by grasses,
chiefly Paspalum vaginatum. It occurs in protected areas of
the seaward side of the lagoon as well as in the landward
side. These U-shaped burrows consist of a broad chamber
with multiple entrances (Figure 7K), inhabited by a huge
amount of C. granulata juveniles and few adults,
characterizing breeding chambers. The strategic location of
these breeding chambers favors the minimum exposure of
the juveniles to the open environment and, consequently, to
predators, and the presence of adults in these galleries suggest
parental care. Decapod breeding chambers have been reported
by Curran (1976), Verde & Martinez (2004) and Lewi &
Goldring (2006) in the fossil recorded, mostly associated with
thalassinidean burrows. Recently, Genise et al. (2008)
reported the occurrence of crayfish breeding burrows in
Cretaceous rocks from Patagonia (Argentina), Cellicalichnus
meniscatus and Dagnichinus titoi, but none of them share
the same morphology observed in the C. granulata breeding
burrows at Peixe Lagoon.
Ant nests are constructed in incipient sandy soils covered
by P. vaginatum and the cactus Cereus sp., in the inner
vegetated areas of the lateral plains and sand bars, and are
completely subterraneous, not showing well-developed
external mounds (Figure 7L). Unfortunately, the effort in
making resin casts to evaluate the nest architecture failed.
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The aeolian dunes suite
This suite is dominated by insect, reptile, bird and mammal
trackways and burrows. Mole cricket burrows and dune
scarab beetles burrows and trackways are the most commonly
observed insect biogenic structures (Figures 4B, 7E, G). Horizontal, irregularly superficial feeding burrows of mole
crickets (Grilotalpidae) are frequently intercepted by bird
trackways. Intersections and bifurcations are common in the
mole cricket burrows, which usually show collapsed roofs
(Figure 7G). Small sandy shore lizard trackways also occur,
exhibiting two rows of alternate diminutive tracks separated
by a medial and continuous sinusoidal groove with discrete
lateral ridges (Figures 7H, I). Mammal biogenic structures are
represented in the aeolian dunes suite by extensive, multichambered gallery systems excavated by small rodents of
the species Ctenomys flamarioni and C. minutus.
The wet interdune suite
This suite is dominated by long, straight to sinuous
shallow horizontal burrows randomly bifurcated or trifurcated
made on sandy softgrounds by mole crickets (Grilotalpide)
in search for food (Figure 7J). The burrow limits are well
defined, but without lining. The burrow roofs are often
PRESERVATION POTENTIAL OF BIOGENIC
SEDIMENTARY STRUCTURES IN THE
PSILONICHNUS ICHNOCOENOSIS
The seaward side of Peixe Lagoon is dynamic setting,
due to the high-energy depositional processes resulting from
streams, storms and winds. For this reason, the potential for
preservation of most of the biogenic structures produced in
this setting is low. Fluctuations in lagoon level may flood the
lagoon edges, destroying partially or totally the Y-shaped
crab burrows commonly present in this setting. The
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
taphonomic result may be the absence of biogenic
sedimentary structures or the preservation of unbranched
vertical shafts, due to the destruction of the upper part of the
burrows by bottom currents. Similar findings were recorded
by Gingras et al. (2000) at Willapa Bay (Washington, USA).
219
Epigenic biogenic sedimentary structures are easily
destroyed in dry environments by wind action, which reworks
the sand grains, erasing the trackways in a few hours. Thus,
arthropod and vertebrate trackways that form the subaerial
ichnocoenosis of Peixe Lagoon have little chance of being
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Figure 7. Biogenic sedimentary structures of the seaward side of Peixe Lagoon. A-C, Vertical burrows of Ocypode quadrata, in beach
stiffgrounds, close to the aeolian dunes (A) and in the base of the aeolian dunes (B), and its cast (C); D, insect horizontal feeding burrow
(?scarab beetle); E, scarab beetle trails departing from vertical burrows; F, mammal trackway; G, mole cricket complex feeding burrow; H,
a looping lizard trackway (from left top to center) going down the sand dune and the moment when the lizard trapped a beetle, going up
after feeding on its prey (from center to right top); I, another meeting between a lizard and a beetle, now in horizontal, indertune sands;
J, mole cricket mosaic horizontal feeding burrows; K, multi-chambered U-shaped nesting burrows of Chasmagnatus granulata with adults
patrolling the burrow entrances; L, single ant nest opening in dry sandy substrates. Biogenic sedimentary structures illustrated in A-D, F
belong to the backshore suite; in E, G-I, to the aeolian dunes suite; in J, to the wet interdune suite; and in K-L, to the vegetated margin suite.
Scale bars: 10 cm.
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REVISTA BRASILEIRA DE PALEONTOLOGIA, 12(3), 2009
preserved in the fossil record. Trackways produced in wet
interdune deposits, otherwise, are generally gently covered
by the sand blown by the wind, having a greater
preservation potential. However, the sands that fill the
trackway depressions have the same composition and
grain size as those forming the trackway-bearing substrate.
The final result is the amalgamation of the sands after bed
compaction, reducing drastically the preservation
potential of the trackways in the fossil record. The
taphonomic response is the little amount of bioturbation
and the low ichnodiversity that is characteristic of the
Psilonichnus Ichnofacies (Pemberton et al., 1992b, 1997,
2001; MacEachern, 2001; Buatois et al., 2002), in contrast
with the more diverse and abundant record observed in
modern Psilonichnus ichnocoenosis (Frey & Pemberton,
1987).
The better chances of preserving tracks and trackways
are in the lagoon edges and shallow bottoms, where crabs
and vertebrates (especially birds) move around looking for
food. The biogenic sedimentary structures produced in
muddy sand substrates are recovered by clean sands blown
by the wind, increasing their preservation potential. The
contrast between these two deposits may be enhanced during
diagenesis, favoring their observation in the fossil record.
The presence of microbial mats also helps in preservation of
tracks and trackways, forming a superficial resistant film that
prevents desiccation and casts perfectly the biogenic
structures as undertracks (Goldring & Seilacher, 1971;
Seilacher, 2008).
The biogenic structures with the highest preservation
potential in the Psilonichnus ichnocoenosis at the Peixe
Lagoon are the Y- and J-shaped burrows that comprise the
lagoon edge suite. The stiffground nature of the substrate
maintains the galleries’ opening for relatively long periods,
favoring their infill by different sediments and highly
increasing their chances of preservation. This pattern is in
accordance with the Psilonichnus Ichnofacies paradigm,
originally discussed by Frey & Pemberton (1987) and not
refuted until now.
The subaquatic ichnocoenosis present in the lagoon floor
(representative of the impoverished Cruziana
Ichnofacies), and the lagoon edge suite of the Psilonichnus
ichnocoenosis succeed each other in the studied area,
creating a tiering marked by the palimpsest imprint of
burrows from both assemblages, in response to water level
fluctuations. Large Y-shaped burrows of C. granulata
(lagoon edge suite) always overprint the tiny Skolithoslike vertical shafts made by polychaetes, which are the
most evident structures of the subaquatic ichnocoenosis
in vertical exposure. Openings of the Y-shaped burrows
are generally preserved, and they are passively filled by
clean, fine-grained, quartz sands showing a faint
lamination. The preservation of the Y morphology and the
characteristics of the infill suggest that erosional processes are rare and sedimentation rates low, allowing inference
of the development of aggradational depositional cycles.
Two types of modern Psilonichnus burrows can be
found in Peixe Lagoon Psilonichnus ichnocoenosis: the
shorter, Y-shaped burrow produced by C. granulata in
backbarrier settings, and the longer, faint Y-shaped burrow
produced by O. quadrata in the beach side of the dune
ridge, in the backshore zone (Figure 10). The average
proportion between the burrow lengths is 1:2 and is linked
to the depth of ground water level: while the freatic level
is established at a depth of 30 cm at the lagoon edges that
comprise the backbarrier deposits, in the backshore areas
it occurs at a depth of around 60 cm. Water table
fluctuations have been reported as an important feature
that controls the distribution of biogenic sedimentary
structures in subaerial continental settings, with the
position of the water table being the major factor governing
the archetypical continental ichnofacies (Buatois &
Mángano, 2004, 2007; Hasiotis, 2007; Netto, 2007; Gibert
& Sáez, 2009). Therefore, the differences between burrow
lengths of Psilonichnus specimens in the same
ichnoassociation may be used to interpret different depths
of the water table level also in marginal marine settings
where the Psilonichnus Ichnofacies occurs, helping to
demarcate autocyclic stratigraphic surfaces.
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STRATIGRAPHIC IMPLICATIONS
EFFECTS OF SEA-LEVEL FLUCTUATIONS
The mix of terrestrial, quasimarine, and marine conditions
of the environmental settings bearing the Psilonichnus
Ichnofacies denotes its transitional character. As a
consequence, the Psilonichnus ichnocoenosis is directly
exposed to eustatic variations and to other phenomena related
to sea level oscillations, which promote episodic, but frequent
erosion of the substrates. For this reason, the preservation
potential of most of the biogenic sedimentary structures that
comprise the Psilonichnus ichnocoenosis is low. In the
Psilonichnus ichnocoenosis at Peixe Lagoon, however,
epigenic biogenic sedimentary structures are abundant and
diverse, suggesting low erosion rates and relatively stable
environmental conditions.
The Coastal Plain of Rio Grande do Sul State (PCRS) is
composed of four distinct parallel, northwest-southeast
oriented marine terraces formed by transgressive-regressive
events, the former three representing the ancient (Pleistocene)
coastlines and the fourth making up the actual coast (Tomazelli
& Villwock, 2000). The Peixe Lagoon area was flooded during
the last transgressive peak (5 ka), and has been experiencing
a shoaling up since them, by the dominance of progradational
processes (Villwock, 1984). The environmental stability
pointed out by the ichnofauna and the recurrent succession
of the ichnocoenoses suggest the maintenance of the facies
distribution pattern since then and allow the inference of the
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
establishment of a stillstand period in the last thousand years.
The equilibrium between the fluvial and marine input allows
the substrate occupation by an impoverished, opportunistic
marine endobenthic invertebrate fauna in the lagoon and a
mixed quasimarine and non marine fauna in the surrounding
settings (Figure 8).
The theoretical analysis of the stratigraphic models
suggests that the sea-level fall, during the establishment of a
lowstand system tract, should enhance the development of
incised valleys and/or channels in the marginal marine zone,
eroding most of the biogenic sedimentary structures
produced during stillstand periods (Taylor & Gawthorpe,
1993; Netto, 2001). The progradation of continental deposits
should favor the development of soils in the area previously
occupied by lagoons, and roots and edaphic insects should
obliterate most of the previous bioturbation that could have
existed. Thus, the potential of preservation of the
Psilonichnus Ichnofacies record under this circumstance is
minimal. A similar situation can be observed in the upper
deposits of the PCRS Barrier III (Upper Pleistocene) cropping
out in Osório (northern littoral of Rio Grande do Sul State)
(Gibert et al., 2006).
On the contrary, transgressive events that occur in
response to eustatic variations are less erosive and can
preserve a good amount of the biogenic sedimentary
structures produced in marginal marine settings, with little
damage to the original morphology (Taylor & Gawthrope,
1993; Netto, 2001). Thus, the Psilonichnus Ichnofacies
record has a greater chance of being preserved during
transgressive system tracts. Considering that the
ichnofacies distribution originally extends to the
backbarrier zone (Frey & Pemberton, 1987), it is possible
to propose that the stratigraphic surface developed right
above the deposits bearing Psilonichnus Ichnofacies
221
could be assumed as a ravinement surface, being, in this
case, the transgressive surface or, in an extreme situation,
the maximum flooding surface (Figure 9). However, if
transgressive surfaces occur associated with a discrete
variation in the sea level base, the complete record of the
Psilonichnus Ichnofacies may be eroded or even obliterated
by the development of the Skolithos Ichnofacies. This may
be the case with Peixe Lagoon, where the Psilonichnus
ichnocoenosis occurs at 60-70 m above sea level, being an
easy target for marine ravinement events. In carbonate
deposits, however, the potential of preservation of the
Psilonichnus ichnocoenosis is higher, as early diagenesis
provides sediment cohesiveness in a few months.
FINAL REMARKS
The analysis of the biogenic sedimentary structures
present on the seaward side of Peixe Lagoon reveals the
existence of an ichnocoenosis with the same
characteristics of the assemblage originally described by
Frey & Pemberton (1987) and which served as model to
define to the Psilonichnus Ichnofacies. In spite of most
ancient occurrences of Psilonichnus Ichnofacies being
related to backshore settings, the Psilonichnus
ichnocoenosis present in the study area reinforces its
extension to backbarrier settings. According to previous
discussion, the preservation of the Psilonichnus
ichnocoenosis of the seaward side of Peixe Lagoon in the
fossil record depends on the establishment of a new
transgressive system tract in the Coastal Plain of the Rio
Grande do Sul State. Once preserved, the biogenic
sedimentary structures of the Psilonichnus ichnocoenosis
may be used as indicators of sea level changes in the PCRS,
like the marine and abrasion terraces currently.
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Figure 8. Schematic drawing showing the distribution of the Psilonichnus ichnocoenosis in the channel mouth area of Peixe Lagoon and
its narrow relationship with the Psilonichnus Ichnofacies.
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Figure 9. Estimated stratigraphic events and their effect on the preservation of the Psilonichnus Ichnofacies. A, actual stillstand period,
in which the biogenic sedimentary structures of the Psilonichnus ichnocoenosis have been made; B, sea-level fall and progressive
erosion of the actual deposits represented in A by progradation of the continental deposits; C, final stage of the hypothetic lowstand
system tract represented in B, in which the Psilonichnus ichnocoenosis record could disappear completely; D, sea-level rise and
consequent marine flooding events recovering the actual deposits represented in A, characterizing a hypothetic transgressive system
tract, which could preserve, rather than destroy, the Psilonichnus ichnocoenosis record.
Figure 10. Schematic profile showing how the freatic water table controls the length of the Y-shaped burrows made in backbarrier (C.
granulata shorter burrows) and backshore (O. quadrata longer burrows) zones.
NETTO & GRANGEIRO – THE PSILONICHNUS ICHNOCOENOSIS AT PEIXE LAGOON, S BRAZIL
ACKNOWLEDGMENTS
The authors thank: IBAMA, through the Peixe Lagoon
National Park administration, for the research license, which
allowed access to the protected areas; CNPq, for research
funds (research grants 304811/2004-1, 474345/2003-3, 303041/
2007-2, and 479457/2007-7); and CAPES, for the graduate
scholarship granted to MEG during the first years of
development of this work. Drawings were originally created
and drawn by MEG. The authors are also grateful to C.H.
Nowatzki, E.L.C. Lavina, and R.C. Lopes, for sedimentologic
discussion during field trips, M.Z. de Oliveira, for helping in
field trips and in electronic edition of figures, and J.M. de
Gibert, F.M.W. Tognoli, and J.R. Garrison, Jr., who critically
reviewed the original manuscript, helping to improve its final
version.
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Received in May, 2009; accepted in November, 2009.